Long-term responses of populations and communities of trees to selective logging in tropical rain forests in Guyana

Since only a small area of Guyana's forest can be effectively protected and because timber harvesting is an important source of income, logged forests will play an important role in the conservation of biodiversity in Guyana. Selective logging, in which only a few trees per hectare are harvested and after which forest remains available, is potentially a good way to utilise the forest without destroying it. In Guyana hard wood from selective logging is an important source of income. As in other tropical countries, sustainable forest management should result in sustained timber yields over long periods of time to provide lasting revenues and to secure livelihoods, while on the other hand also diversity should be conserved as much as possible. To be able to define criteria for sustainable forest management, information on the long-term effects of logging is needed. Selective logging creates openings in the forest canopy, which results in increased light availability in the forest understorey. As a consequence of this increased light availability some tree species (the pioneers) are able to grow much faster. On the long term this may result in changes in species composition of the forest. The aim of the investigations described in this thesis was to determine the long-term effects of logging on tree population dynamics, forest composition and tree diversity and to evaluate the sustainability of alternative forest management scenarios for both future timber yields and biodiversity conservation. To investigate these long-term effects, a field study was done in logged and non-logged forests in Guyana and additionally a forest simulation model was developed to evaluate different management scenarios. This population dynamics model simulates growth, mortality and recruitment of trees and makes projections of forest composition and available hard wood in the course of decades. The results of the field study showed that increased light availability after logging is especially advantageous for pioneer species. The abundance of inherently slow growing tree species decreased, but recovered again in the course of years after logging. Model simulations showed, however, that selective logging did not severely affect forest composition. Even in simulations of the most intensive way of logging (12 trees ha-1, every 25 years) forest composition remained rather intact. This is probably due to the fact that in forests in central Guyana, pioneer species are very rare and thus will not easily dominate the forest after logging. After logging once using high harvest intensities of 12 trees ha-1, it took, however, more than 100 years before harvestable timber volumes were comparable again with a baseline (non-logging) situation. For slow growing tree species it even took more than 160 years after logging before the abundance of stems was comparable again with the baseline situation. Projected recovery periods were, however, substantially longer than the currently in Guyana advised length of felling cycle of 60 years. Highest total timber yields were achieved if trees were harvested every 25 years using high harvest intensities. At the same time this approach also resulted in a fast depletion of the available commercial timber volumes in the forest and thus reduced timber yields. The results of the investigations in this thesis can be used to determine criteria for sustainable forest management in Guyana.

[1]  R. Boot,et al.  Approaches to developing sustainable extraction systems for tropical forest products , 1995 .

[2]  J. Connell Diversity in tropical rain forests and coral reefs. , 1978, Science.

[3]  M. Swaine The ecology of tropical forest tree seedlings , 1996 .

[4]  D. Lieberman,et al.  Forest tree growth and dynamics at La Selva, Costa Rica (1969-1982) , 1987, Journal of Tropical Ecology.

[5]  P. Köhler,et al.  Modelling anthropogenic impacts on the growth of tropical rain forests , 2000 .

[6]  D. Sheil Long-term observations of rain forest succession, tree diversity and responses to disturbance. , 2001, Plant Ecology.

[7]  L. Poorter Light-dependent changes in biomass allocation and their importance for growth of rain forest tree species , 2001 .

[8]  Stephen J. Wright,et al.  Light-Gap disturbances, recruitment limitation, and tree diversity in a neotropical forest , 1999, Science.

[9]  R. Kobe LIGHT GRADIENT PARTITIONING AMONG TROPICAL TREE SPECIES THROUGH DIFFERENTIAL SEEDLING MORTALITY AND GROWTH , 1999 .

[10]  P. R. van Gardingen Growth and yield modelling: applications of SYMFOR to evaluate silvilcultural systems , 1999 .

[11]  Annabel Porté,et al.  Modelling mixed forest growth: a review of models for forest management , 2002 .

[12]  Jean Gérard,et al.  Major timber trees of Guyana: timber characteristics and utilization. , 1996 .

[13]  R. Condit,et al.  Inferred longevity of Amazonian rainforest trees based on a long-term demographic study , 2004 .

[14]  D. Burslem,et al.  The interpretation and misinterpretation of mortality rate measures , 1995 .

[15]  O. Dam Forest filled with gaps : effects of gap size on water and nutrient cycling in tropical rain forest : a study in Guyana , 2001 .

[16]  D. Lieberman,et al.  Growth rates and age-size relationships of tropical wet forest trees in Costa Rica , 1985, Journal of Tropical Ecology.

[17]  Bette A. Loiselle,et al.  SEED AND SEEDLING ECOLOGY OF NEOTROPICAL MELASTOMATACEAE , 1993 .

[18]  T. Lovejoy,et al.  Rain-forest fragmentation and the phenology of Amazonian tree communities , 2003, Journal of Tropical Ecology.

[19]  Cannon,et al.  Tree species diversity in commercially logged bornean rainforest , 1998, Science.

[20]  E. Álvarez-Buylla,et al.  Demography and allometry of Cecropia obtusifolia, a neotropical pioneer tree - an evaluation of the climax-pioneer paradigm for tropical rain forests , 1992 .

[21]  David F. R. P. Burslem,et al.  Disturbing hypotheses in tropical forests , 2003 .

[22]  Miguel Martínez-Ramos,et al.  The consequences of crown traits for the growth and survival of tree saplings in a Mexican lowland rainforest , 2003 .

[23]  Josefina Barajas-Morales Wood Specific Gravity in Species from Two Tropical Forests in Mexico , 1987 .

[24]  P. Savill,et al.  Long-term permanent plot observations of vegetation dynamics in Budongo, a Ugandan rain forest , 2000, Journal of Tropical Ecology.

[25]  M. Swaine,et al.  On the definition of ecological species groups in tropical rain forests , 1988, Vegetatio.

[26]  Andreas Huth,et al.  Sustainable timber harvesting in Venezuela: a modelling approach , 2001 .

[27]  Daniel Sabatier,et al.  An analysis of the floristic composition and diversity of Amazonian forests including those of the Guiana Shield , 2000, Journal of Tropical Ecology.

[28]  E. Suzuki Diversity in specific gravity and water content of wood among Bornean tropical rainforest trees , 1999, Ecological Research.

[29]  J. E. Winstead Fiber Tracheid Length and Wood Specific Gravity of Seedlings as Ecotypic Characters in Liquidambar Styraciflua L. , 1972 .

[30]  M. Werger,et al.  Regeneration of timber trees in a logged tropical forest in North Bolivia , 2004 .

[31]  M. Peña-Claros Changes in Forest Structure and Species Composition during Secondary Forest Succession in the Bolivian Amazon1 , 2003 .

[32]  A. Gentry,et al.  Changes in Plant Community Diversity and Floristic Composition on Environmental and Geographical Gradients , 1988 .

[33]  R. Ek Botanical diversity in the tropical rain forest of Guyana. , 1997 .

[34]  Susan A. Foster,et al.  The relationship between seed size and establishment conditions in tropical woody plants , 1985 .

[35]  G. B. Williamson Pattern and Seral Composition in an Old‐growth Beech‐Maple Forest , 1975 .

[36]  Bai-Lian Li,et al.  Habitat Destruction and the Extinction Debt Revisited , 1996 .

[37]  Frans Bongers,et al.  The ecology of lianas and their role in forests , 2002 .

[38]  Boris Zeide,et al.  Analysis of Growth Equations , 1993 .

[39]  E. Wilson,et al.  The Diversity of Life , 1993, Politics and the Life Sciences.

[40]  D. Burslem,et al.  Species diversity, susceptibility to disturbance and tree population dynamics in tropical rain forest , 1999 .

[41]  T. Givnish On the causes of gradients in tropical tree diversity , 1999 .

[42]  D. S. Hammond,et al.  CHARACTER CONVERGENCE, DIVERSITY, AND DISTURBANCE IN TROPICAL RAIN FOREST IN GUYANA , 2001 .

[43]  B. Finegan Pattern and process in neotropical secondary rain forests: the first 100 years of succession. , 1996, Trends in ecology & evolution.

[44]  P. R. van Gardingen,et al.  Grouping tree species for analysis of forest data in Kalimantan (Indonesian Borneo) , 2002 .

[45]  Oscar García,et al.  Evaluating forest Growth Models , 1997 .

[46]  Jerome K. Vanclay,et al.  GROWTH AND YIELD OF A TROPICAL RAIN FOREST IN THE BRAZILIAN AMAZON 13 YEARS AFTER LOGGING , 1995 .

[47]  D. Clark,et al.  Age and Long‐term Growth of Trees in an Old‐growth Tropical Rain Forest, Based on Analyses of Tree Rings and 14C1 , 2003 .

[48]  E. Webb,et al.  Tree community diversity of lowland swamp forest in Northeast Costa Rica, and changes associated with controlled selective logging , 1998, Biodiversity & Conservation.

[49]  William H. Press,et al.  Book-Review - Numerical Recipes in Pascal - the Art of Scientific Computing , 1989 .

[50]  K. Ickes,et al.  Resprouting of woody saplings following stem snap by wild pigs in a Malaysian rain forest , 2003 .

[51]  O. Phillips,et al.  Dynamics and species richness of tropical rain forests. , 1994, Proceedings of the National Academy of Sciences of the United States of America.

[52]  L. Poorter,et al.  Light environment and tree strategies in a Bolivian tropical moist forest: an evaluation of the light partitioning hypothesis , 2003, Plant Ecology.

[53]  P. Grubb Rainforest dynamics: the need for new paradigms , 1996 .

[54]  M. Nowak,et al.  Habitat destruction and the extinction debt , 1994, Nature.

[55]  F. Bongers,et al.  The effect of canopy gaps on growth and morphology of seedlings of rain forest species , 1988, Oecologia.

[56]  R. Zagt Tree demography in the tropical rain forest of Guyana. , 1997 .

[57]  S L Lewis,et al.  Pattern and process in Amazon tree turnover, 1976-2001. , 2004, Philosophical transactions of the Royal Society of London. Series B, Biological sciences.

[58]  Changhui Peng,et al.  Growth and yield models for uneven-aged stands: past, present and future , 2000 .

[59]  R. Primack,et al.  Growth Rates and Population Structure of Moraceae Trees in Sarawak, East Malaysia , 1985 .

[60]  Christopher Uhl,et al.  Logging damage during planned and unplanned logging operations in the eastern Amazon , 1996 .

[61]  N. Brokaw,et al.  Niche versus chance and tree diversity in forest gaps. , 2000, Trends in ecology & evolution.

[62]  Hendrik Poorter,et al.  Inherent variation in plant growth : physiological mechanisms and ecological consequences , 1998 .

[63]  L. Poorter,et al.  Growth and carbon partitioning of tropical tree seedlings in contrasting light environments. , 1998 .

[64]  S. Pacala,et al.  Forest models defined by field measurements : Estimation, error analysis and dynamics , 1996 .

[65]  J. Chave Study of structural, successional and spatial patterns in tropical rain forests using TROLL, a spatially explicit forest model , 1999 .

[66]  Yadvinder Malhi,et al.  Tropical forests and global atmospheric change: a synthesis. , 2004, Philosophical transactions of the Royal Society of London. Series B, Biological sciences.

[67]  D. Lieberman,et al.  Forests are Not Just Swiss Cheese: Canopy Stereogeometry of Non‐Gaps in Tropical Forests , 1989 .

[68]  Robin L. Chazdon,et al.  SPATIAL HETEROGENEITY OF LIGHT AND WOODY SEEDLING REGENERATION IN TROPICAL WET FORESTS , 1999 .

[69]  L. Poorter,et al.  The importance of seed mass for early regeneration in tropical forest: a review , 2003 .

[70]  D. S. Hammond,et al.  Seed Size of Woody Plants in Relation to Disturbance, Dispersal, Soil Type in Wet Neotropical Forests , 1995 .

[71]  V. Favrichon Classification des espèces arborées en groupes fonctionnels en vue de la réalisation d'un modèle de dynamique de peuplement en forêt Guyanaise , 1994 .

[72]  P. R. van Gardingen,et al.  An individual-based spatially explicit simulation model for strategic forest management planning in the eastern Amazon , 2004 .

[73]  Robin L. Chazdon,et al.  PHOTOSYNTHETIC LIGHT ENVIRONMENTS IN A LOWLAND TROPICAL RAIN FOREST IN COSTA RICA , 1984 .

[74]  A. V. Kekem,et al.  Soils of the rainforest in central Guyana. , 1996 .

[75]  Douglas W. Yu,et al.  Demographic Threats to the Sustainability of Brazil Nut Exploitation , 2003, Science.

[76]  D. Lieberman,et al.  Simulation of Growth Curves from Periodic Increment Data , 1985 .

[77]  C. Augspurger Light requirements of neotropical tree seedlings: a comparative study of growth and survival , 1984 .

[78]  I. Válio,et al.  Seed Size, Seed Germination, and Seedling Survival of Brazilian Tropical Tree Species Differing in Successional Status1 , 2001 .

[79]  P. Gardingen,et al.  The SYMFOR framework for individual-based spatial ecological and silvicultural forest models , 2001 .

[80]  Effects of selective logging on tree diversity, composition and plant functional type patterns in a Bornean rain forest , 2003 .

[81]  William H. Press,et al.  Numerical recipes in C. The art of scientific computing , 1987 .

[82]  H. Prins,et al.  Gap-size niche differentiation by tropical rainforest trees: a testable hypothesis or a broken-down bandwagon? , 1998 .

[83]  R. Fisher,et al.  The Relation Between the Number of Species and the Number of Individuals in a Random Sample of an Animal Population , 1943 .

[84]  F. Putz Biodiversity Conservation in the Context of Tropical Forest Management , 2000 .

[85]  S. Gourlet-Fleury Indices de compétition en forêt dense tropicale humide : étude de cas sur le dispositif sylvicole expérimental de Paracou (Guyane française) , 1998 .

[86]  David B. Clark,et al.  LIFE HISTORY DIVERSITY OF CANOPY AND EMERGENT TREES IN A NEOTROPICAL RAIN FOREST , 1992 .

[87]  N. Brokaw Gap-Phase Regeneration of Three Pioneer Tree Species in a Tropical Forest , 1987 .

[88]  C. Uhl,et al.  Vegetation Dynamics in Amazonian Treefall Gaps , 1988 .

[89]  F. Bongers,et al.  Formation and closure of canopy gaps in the rain forest at Nouragues, French Guiana , 1996, Vegetatio.

[90]  N. Picard,et al.  A comparison of five indirect methods for characterizing the light environment in a tropical forest , 2001 .

[91]  A. Newton Dynamics of Tropical Communities , 1999 .

[92]  Lourens Poorter,et al.  Growth responses of 15 rain‐forest tree species to a light gradient: the relative importance of morphological and physiological traits , 1999 .

[93]  A. Di Fiore,et al.  Increasing biomass in Amazonian forest plots. , 2004, Philosophical transactions of the Royal Society of London. Series B, Biological sciences.

[94]  S. Hubbell,et al.  The demographics of resprouting in tree and shrub species of a moist tropical forest , 2000 .

[95]  H. Balslev,et al.  Growth rates and mortality patterns of tropical lowland tree species and the relation to forest structure in Amazonian Ecuador , 1994, Journal of Tropical Ecology.

[96]  S. Davies TREE MORTALITY AND GROWTH IN 11 SYMPATRIC MACARANGA SPECIES IN BORNEO , 2001 .

[97]  Plinio Sist,et al.  An experimental comparison of different harvesting intensities with reduced-impact and conventional logging in East Kalimantan, Indonesia , 1997 .

[98]  P. R. van Gardingen,et al.  The SYMFOR framework for modelling the effects of silviculture on the growth and yield of tropical forests , 2001 .

[99]  Jianguo Liu,et al.  Individual-based simulation models for forest succession and management , 1995 .

[100]  Stephen P. Hubbell,et al.  Sapling Survival, Growth, and Recruitment: Relationship to Canopy Height in a Neotropical Forest , 1991 .

[101]  D. Lieberman,et al.  Canopy closure and the distribution of tropical forest tree species at La Selva, Costa Rica , 1995, Journal of Tropical Ecology.

[102]  S. Levin,et al.  Scaling from Trees to Forests: Analysis of a Complex Simulation Model , 1997 .

[103]  Daniel Sabatier,et al.  A spatial model of tree α-diversity and tree density for the Amazon , 2003, Biodiversity & Conservation.

[104]  A. D. Johns Effects of "selective" timber extraction on rain forest structure and composition and some consequences for frugivores and folivores , 1988 .

[105]  J. Terborgh,et al.  Concerted changes in tropical forest structure and dynamics: evidence from 50 South American long-term plots. , 2004, Philosophical transactions of the Royal Society of London. Series B, Biological sciences.

[106]  P. R. van Gardingen,et al.  Financial and ecological analysis of management options for logged-over Dipterocarp forests in Indonesian Borneo , 2003 .

[107]  Daniel Sabatier,et al.  Tree Diversity in Tropical Rain Forests: A Validation of the Intermediate Disturbance Hypothesis , 2001, Science.

[108]  N. Brokaw Gap-phase regeneration in a tropical forest. , 1985 .

[109]  F. Houllier,et al.  Ecological meaning and predictive ability of social status, vigour and competition indices in a tropical rain forest (India) , 1999 .

[110]  L. H. V. Ulft The effect of seed mass and gap size on seed fate of tropical rain forest tree species in guyana. , 2004 .

[111]  Stuart J. Davies,et al.  Comparative ecology of 11 sympatric species of Macaranga in Borneo: tree distribution in relation to horizontal and vertical resource heterogeneity , 1998 .

[112]  Andreas Huth,et al.  Concepts for the aggregation of tropical tree species into functional types and the application to Sabah's lowland rain forests , 2000, Journal of Tropical Ecology.

[113]  Jerome K. Vanclay,et al.  Modelling Forest Growth and Yield: Applications to Mixed Tropical Forests , 1994 .

[114]  R. Bunce,et al.  A Long-term Surveillance System for British Woodland Vegetation , 1978 .

[115]  F. Putz,et al.  BIOLOGICAL CHALLENGES FOR CERTIFICATION OF TROPICAL TIMBER , 1996 .

[116]  P. Sist,et al.  Harvesting intensity versus sustainability in Indonesia , 1998 .

[117]  D. L. Scarnecchia,et al.  Fundamentals of Ecological Modelling , 1995 .

[118]  S. Hubbell,et al.  SPECIES-AREA AND SPECIES-INDIVIDUAL RELATIONSHIPS FOR TROPICAL TREES : A COMPARISON OF THREE 50-HA PLOTS , 1996 .

[119]  H. Steege,et al.  Long-term effect of timber harvesting in the Bartica Triangle, Central Guyana. , 2002 .

[120]  R. Condit,et al.  Pervasive alteration of tree communities in undisturbed Amazonian forests , 2004, Nature.

[121]  P. Hout Reduced Impact Logging in the Tropical Rain Forest of Guyana: Ecological, Economic and Silvicultural Consequences, Tropenbos Guyana Series 6 , 1999 .

[122]  M. Peña-Claros Secondary forest succession : processes affecting the regeneration of Bolivian tree species , 2001 .

[123]  D. Sheil,et al.  Tree Species Diversity in Logged Rainforests , 1999, Science.

[124]  N. Manokaran,et al.  Recruitment, growth and mortality of tree species in a lowland dipterocarp forest in Peninsular Malaysia , 1987, Journal of Tropical Ecology.

[125]  H. S. Horn,et al.  The Ecology of Secondary Succession , 1974 .

[126]  P. Ashton,et al.  Phenology and fecundity in 11 sympatric pioneer species of Macaranga (Euphorbiaceae)in Borneo. , 1999, American journal of botany.

[127]  Jianguo Liu,et al.  FORMOSAIC: an individual-based spatially explicit model for simulating forest dynamics in landscape mosaics , 1998 .

[128]  R. Verburg,et al.  Temporal changes in species diversity, composition, and plant functional types after logging in a Bornean rain forest. , 2003 .

[129]  F. Putz,et al.  Effects of lianas on growth and regeneration of Prioria copaifera in Darien, Panama , 2004 .

[130]  S. Hubbell,et al.  A spatially explicit model of sapling growth in a tropical forest: does the identity of neighbours matter? , 2004 .

[131]  Burslem,et al.  Biotic Interactions in the Tropics , 2005 .

[132]  S. Thomas Asymptotic height as a predictor of growth and allometric characteristics in malaysian rain forest trees , 1996 .

[133]  P. R. van Gardingen,et al.  An individual-based spatially explicit tree growth model for forests in East Kalimantan (Indonesian Borneo) , 2003 .

[134]  H. Steege Plant diversity in Guyana: with recommendations for a national protected area strategy. , 2000 .

[135]  William F. Laurance,et al.  RAIN FOREST FRAGMENTATION AND THE DYNAMICS OF AMAZONIAN TREE COMMUNITIES , 1998 .

[136]  J. Chambers,et al.  Ancient trees in Amazonia , 1998, Nature.

[137]  D. Bebber,et al.  Growth and survivorship of dipterocarp seedlings: differences in shade persistence create a special case of dispersal limitation. , 1999, Philosophical transactions of the Royal Society of London. Series B, Biological sciences.

[138]  M. Werger,et al.  Gap-dependence and leaf characteristics of trees in a tropical lowland rain forest in Mexico. , 1992 .

[139]  L. H. van Ulft Regeneration in natural and logged tropical rain forest : modelling seed dispersal and regeneration , 2004 .

[140]  M. Rosenzweig,et al.  Species Diversity in Space and Time , 1995 .

[141]  D. S. Hammond,et al.  Ecology and logging in a tropical rain forest in Guyana: with recommendations for forest management. , 1996 .

[142]  Yadvinder Malhi,et al.  Fingerprinting the impacts of global change on tropical forests. , 2004, Philosophical transactions of the Royal Society of London. Series B, Biological sciences.

[143]  N. Higuchi,et al.  Logging activity and tree regeneration in an Amazonian forest , 1999 .

[144]  Steven F. Oberbauer,et al.  Landscape scale evaluation of understory light and canopy structure : methods and application in a neotropical lowland rain forest , 1996 .

[145]  A. Gentry Tree species richness of upper Amazonian forests. , 1988, Proceedings of the National Academy of Sciences of the United States of America.